Bird Paper One

BY: Bob Montgomerie, Queen’s University | 10 December 2018

When we were writing our Ten Thousand Birds book on the history of ornithology since Darwin, we thought it might be interesting to try to illustrate the growth of the field since the mid-1800s. To do that, we prepared a graph showing the number of articles and books published per year for every fifth year since 1865, using both Zoological Record and, for recent years, Google Scholar. The results were staggering [1], showing an explosive growth in publications on—and presumably knowledge about—birds since the second world war. Since the year 2000, there have been more articles and books published about birds than in the entire period from the beginning of scientific publishing in 1665 until 2000. We can estimate the number of publications before 1865 with some confidence as there were very few bird papers published before that date. The world’s major bird journals did not even start publishing until the mid-to-late 1800s [2].

Untitled

When we compiled that graph, we wondered when the first-ever scientific paper had been published on birds. It had to be after 1664, as the first ever scientific journals [3], Journal des sçavans and Philosophical Transactions, began publishing early in 1665.

To find that first bird paper, I scoured the early issues of both journals, looking at each issue as there was no Zoological Record or Google Scholar coverage that far back. The early issues of Journal des sçavans were devoted largely to obituaries, astronomy, and Cartesian philosophy, and Philosophical Transactions focused mainly on optics, astronomy, and other physical phenomena in its earliest years, though most issues had at least one paper on a biological/medical topic [4].

Bee_hummingbird_(Mellisuga_helenae)_adult_male_in_flight
Male Bee Hummingbird

Although birds were mentioned in several papers in the first few years of scientific publication, the first paper exclusively about birds did not appear until May 1693—in the 17th volume, and 200th issue, of Philosophical Transactions. That paper was attributed to the noted English botanist Nehemiah Grew [5] who published a letter (by a Mr Hamersly [6]) describing a hummingbird. He called the bird both ‘Hum Bird’ and ‘Tomineius’, the latter a Spanish word derived from ‘tomino‘ which was a measure of weight equal to 12 grains (0.78 g). In his Ornithology of Francis Willughby, published in 1678, John Ray suggests that the name ‘Tomineius‘ reflects the weight of the bird. But the smallest hummingbird—the Bee Hummingbird (Mellisuga helenae) of Cuba—weighs three times that. I suspect that the name ‘Tomineius’ was just meant to indicate that the bird was extremely small.

Antillean-Crested-Hummingbird-Orthorhyncus-cristatusWe don’t know for sure which species Hamersly was referring to as tomineius was probably a general term for all hummingbird species. In his 1590  book on the West Indies, for example, José de Acosta says that hummingbirds were called ‘Tomineios‘ in Peru.

A hand-written annotation  in some manuscript notes [7], presumably by Hamersly, found in a copy of Richard Ligon’s 1657 book on Barbados says that “I sent this description of this bird to doctor Grew one of the Royal Society & he caused it to be printed in their philosophical transactions. This suggests that Hamersly was describing one of the 3 species that are common on Barbados. By ‘this bird’, the annotator was referring to the picture below right, which, though crude, looks most like the Green-throated Carib (Eulampis holosericeus. By its size, however, I think Hammersly must have been referring to the Antillean Crested Hummingbird (Orthorhyncus cristatus) which weighs about 3 grams, the smallest hummingbird on the island. It’s too bad that we don’t really know who Hamersly was, nor which species he was describing.

GtCarib
Green-throated Carib (L), and hummingbird illustration (R) from manuscript notes [7]

Grew, or rather Hamersly, made a number of perceptive observations of the hummingbirds, though recent research has shown that he was not quite correct. Here are a few of the interesting things that Hamersly noted, with comments and what we now know in square brackets:

  • “He is of a most excellent shining green Color…resemble some of our English Drake-heads” [true, both are iridescent green]
  • whole weight was the tenth part of an ounce Avoirdupoise” [this would be about 2.8 g which is about right for the Antillean Crested Hummingbird]
  • They feed by thrusting their Bill and Tongue into the Blossoms of Trees, and so suck the sweet juice of Honey from them” [hummingbirds don’t suck [8]; they take up nectar into their grooved tongue and force it back into their throat by pressing their tongue with their bill as it retracts]
  • I did observe them several years but never heard them sing” [he claims they don’t sing, but they do, as do all of the 50 or species that I know reasonably well. He may have meant they don’t sing a song that sounds like most of the passerine bird songs and that is generally correct]
  • He is called the Hum-bird or Humming Bird because some say he makes a noise like a Spinning Wheel when he flies..I never heard any Noise; besides their Body and Wings are too small to strike the Air to make any Noise” [he is mistaken here, of course, but he later acknowledges that other people have heard them humming. He should have known that mosquitoes make noise so that his comment about size must be wrong.]

He did correctly note that they are very solitary, and suggested that with such a beautiful plumage they may not need to sing well: “so I think this Bird is so beautiful to the Eye, as not at all to please the Ear“. Indeed, recent studies have found such a tradeoff between selection for elaborate song or bright plumage in different groups of bird [9].

In Nehemiah Grew’s day, anyone interested in the sciences could read everything published in all (both) of the scientific journals. Even when I was a PhD student, in the 1970s, it was possible (and de rigueur) to read most of the papers in ecology and evolution published in the major journals, and to read all of the recent papers published on your study organism. Those days are over and few scientists can manage to even be aware of all of the research relevant to their own studies. No wonder many scientists get most of their information about recent studies in their field from Twitter.

Even if you wanted to keep up with research on birds since 2000, you would face a daunting task. The Web of Science [10] says that 127,000 papers have been published on birds from 2000 to 2018. The following graphic shows the distribution of 115,000 of those papers in the best-studied topics:

ornithology
Papers on birds published 2000-2018 on the 10 most common topics

Even focussing on hummingbirds, you would have to read 2383 papers to be fully informed about research published since 2000 (see below). Contrast this to the 36 papers on hummingbird ecology and evolution published during my PhD years, and the 48 papers published on those topics from 1900 until the year I graduated in 1979. We have come along way since Grew began the scientific publications about birds.

hbird papers
Papers on hummingbirds published from 2000-2018 on the 10 most common topics

SOURCES

  • Badyaev AV, Hill GE, Weckworth BV (2002) Species divergence in sexually selected traits: increase in song elaboration is related to decrease in plumage ornamentation in finches. Evolution 56: 412–419

  • Birkhead TR, Wimpenny J, Montgomerie R (2014) Ten Thousand Birds: Ornithology since Darwin. Princeton, NJ: Princeton University Press.

  • Boyle (1865) A way of preserving birds taken out of the egge, and other small fætus’s. Philosophical Transactions 1: 199-201

  • de Acosta J (1590) Historia Natural y Moral de las Indias. Sevilla: Juan de Leon.
  • Grew N (1693a) The description of the American tomineius, or hummingbird. Philosophical Transactions 17: 760-761
  • Grew N (1693b) A query put by Dr. N. Grew, concerning the food of the Humming Bird ; occasioned by the description of it in the transactions. Numb. 200. Philosophical Transactions 17: 815
  • Lefanu W (1971) The Versatile Nehemiah Grew. Proceedings of the American Philosophical Society 115: 502-506
  • Ligon R (1657) A true & exact history of the island of Barbados: Illustrated with a mapp of the island, as also the principall trees and plants there, set forth in their due proportions and shapes, drawne out by their severall and respective scales. Together with the ingenio that makes the sugar, with the plots of the severall houses, roomes, and other places, that are used in the whole processe of sugar-making; viz. the grinding-room, the boyling-room, the filling-room, the curing-house, still-house, and furnaces; all cut in copper. London: Printed for Humphrey Moseley, at the Prince’s Arms in St. Paul’s Church-Yard.

  • Price DJS (1963) Little science, big science. New York: Columbia University Press.

  • Ray J (1678) The Ornithology of Francis Willughby. London: John Martyn.

  • Rico-Guevara A, Fan T-H, Rubega MA (2015) Hummingbird tongues are elastic micropumps. Proceedings of the Royal Society B: Biological Sciences 282:20151014.

  • Shutler D, Weatherhead PJ (1990) Targets of sexual selection: song and plumage of wood warblers. Evolution 44:1967–1977.


Footnotes

  1. staggering results: while the numbers are high, the pattern is typical of all sciences, as described by Derek da Solla Price in his 1963 book
  2. first major ornithological journals: see previous posts, here, here, and here
  3. first ever scientific journals: Journal des sçavans began publishing on 5 Jan 1665, and Philosophical Transactions (later called Philosophical Transactions of the Royal Society) on 6 March of that same year.
  4. one paper on biological topics: Philosophical Transactions did have early papers on snake behaviour, breeding silkworms, and various medical anomalies, for example. There was even an 1865 paper on preserving bird embryos (Boyle 1865) but I don’t really count that as being about birds
  5. Nehemia Grew: was an early Fellow of the Royal Society, and was both editor of Philosophical Transactions (1678-79) and secretary of the RS. He is often called the Father of Plant Anatomy
  6. Mr Hamersly: although Grew published this letter under his own name, he revealed in Grew (1693b) that it was actually written by a Mr Hamersly of Coventry. See here for more information, suggesting it might have been John Hamersly, also referred to here as John Hammersley.
  7. manuscript notes: see here for details
  8. hummingbirds don’t suck: I discovered this tin the 1970s when doing my PhD on hummingbirds but I am sure that all of the other biologists I knew who were studying hummingbirds in those days—Bill Calder, Peter Feinsinger, Lee Gass, Larry Wolf, Reed Hainsworth and Frank Gill—knew this too; Rico-Guevera et al. (2015) recently described the details and physics of this process
  9. tradeoff between song and plumage: see for example, Shutler and Weatherhead (1990) and Badyaev et al. (2002)
  10. Web of Science: These numbers are smaller than what we show in the graph at the top of this essay, because Web of Science focuses only on publications in scientific journals, whereas we graphed all publications about birds. The data from Web of Science show the same patterns as in that graph but only about 7000 bird papers published in 2010, for example. To generate those data I entered bird* and ornithology as topics for the first graph and hummingbird* for the second and searched through the years in question

IMAGES: top graph from Birkhead et al. (2014), drawn by the author; Bee Hummingbird from Wikipedia; stamp from a stamp collection website; Green-thoated Carib (on Barbados) photo from Wikipedia, painting from manuscript notes [7]; bottom two graphs from Web of Science (accessed 9 Dec 2018)

The Sacred Sacred Ibis [reposted]

BY: Bob Montgomerie, Queen’s University | 15 October 2018

446px-Thoout,_Thoth_Deux_fois_Grand,_le_Second_Hermés,_N372.2A
Thoth and ibises

The ancient Greeks usually depicted Thoth—their god of writing, wisdom and magic—as having the head of a bird with a long, down-curving bill.  Until the 1800s, Europeans thought that this bird was probably a curlew, a stork or a heron. Linnaeus believed that the bird must be the Cattle Egret which he called Ardea ibis in the 1758 edition of his Systema Naturae. It was not until the turn of the 19th century that a small group of French scientists and naturalists finally confirmed the connection between Thoth and the head of the African Sacred Ibis (Threskiornis aethiopicus). This species was not unambiguously described until 1790 [1], but it took Napoleon Bonaparte’s military campaign to Egypt to provide the connection between this species and Thoth.

We now know that the sacred ibis was considered (and revered) by the Egyptians to be the earthly form of Thoth. For about a millennium starting in 1100 BC, ibises were frequently mummified as an offering to Thoth, believing that mummification would put the birds on a direct line to the afterlife. As a result, several million sacred ibises were killed, gutted, embalmed and folded with the bill tucked between the tail feathers. The carcasses were then wrapped with linen dipped in resin, and inserted individually or in pairs into urns that were placed in vast underground caverns in cities all along the Nile. Many of these mummified ibises have grains, snakes, snails and other foods in their body cavities, possibly to provide the birds with some food in the afterlife.

But why ibises, and where did all of these birds come from? There can be no doubt that the sacred ibis was a reasonably common bird [2] in swampy areas all along the Nile in the Late and Ptolomeic Periods of ancient Egyptian civilization [3]. Those birds were of great value to nearby villages as they ate the snails that infested fish ponds, snails that harboured parasites dangerous to humans. They were also claimed to feed on flying snakes (?) and generally consumed all kinds of human refuse [4]. No wonder they were considered to be sacred.

At several sites of ancient cities along the Nile, archaeologists have found incredible numbers of mummified ibises: 1.75 million at Saqqara, 4 million at Tuna el-Gebel, for example. Even over a period of 500 years that is a lot of birds per year, likely magnitudes more than could have been hunted in the local marshes for any sustained period. Because of their religious importance, sanctuaries dedicated to the ibis sprang up all over the country, where birds were bred and raised in captivity, processing as many as 20,000 ibises per year for the votive ibis industry. Priests apparently gathered eggs for artificial incubation and tended the large flocks, as well as engaging in a large pottery industry to make urns for the mummified birds. These ibiotropheia may well be the earliest examples of bird-farming that did not involve some form of fowl.

The vast stores of ibis mummies in Egypt were brought to light by Geoffery Saint-Hilaire and Jules-César Savigny, two of the 167 savants [5] who accompanied Napoleon Bonaparte to Egypt from 1798-1801. Savigny noticed that the ibis often appeared in hieroglyphics and tomb paintings, and reasoned that this bird was important to Egyptian culture. He wrote up his discoveries in 1805 as Histoire naturelle et mythologique de l’ibis which included some very nice illustrations.

Savigny
from Savigny (1805) hand-coloured by Louis Bouqet

Georges Cuvier, one of the leading French biologists of the day, was asked by Napoleon to join the Egyptian contingent, but he suggested that Savigny go instead, so he could continue his work on molluscs. But it was Cuvier who first measured two mummified birds brought back from Egypt by Col. Jacques François-Louis Grobert [6] from the catacombs at Saqqara.  Cuvier initially concluded that those birds were probably curlews as they were smaller than some contemporary ibis specimens [7]. He later measured two mummies that Geoffrey Saint-Hilaire had brought home from Thebes. Those measurements plus the colours of some intact feathers convinced him that the mummies collected by Saint-Hilaire were indeed sacred ibises, and his 1804 paper has a very nice summary of his reasoning and all of the previous mis-identifications. Cuvier’s assistant even pieced together the bones from different mummies to make a complete skeleton (see picture below).

Even though the ibis mummies from Saint-Hilaire were not exactly the same size as contemporary birds, Cuvier also used those measurements to bolster his arguments of the fixity of species—evidence that species were created once by a deity and did not change through time. This argument put him at odds with his colleague Lamarck who argued that species changed through geological time.

I was made aware of this ibis story in a new essay in PLoS Biology [8], by Caitlin Curtis, Craig Millar and David Lambert. As Jerry Coyne noted in an essay on his Why Evolution is True site, not many evolutionary biologists seem to be aware of this as an early test of evolutionary change. The reason, I think, is that it was not actually a test [9]. The story is actually rather well known and has been published many times in scientific journals and the popular press ever since Cuvier’s initial publications [10]. While the new essay summarizes many aspects of this story the authors present no evidence in support of some of their claims and I am not entirely convinced by some of their assertions.

CuvierIBIS
Cuvier (1804) identifies the mummies as sacred ibises

When interpreting the past here is always a danger of applying present knowledge and values incorrectly. In this case, I cannot yet tell if my different interpretation of this interesting story is correct. I will need to read the work of Cuvier, Lamarck, Saint-Hilaire and Savigny in the original French and Latin to put the whole story in context but that will take a while, even though all of the relevant texts are now available online. I will revisit the topic when I have done the necessary research.

Whether the details in this new essay by Curtis and colleagues are correctly interpreted or not, it does end with a curious conclusion that I feel deserves some further discussion: Of great importance is the reminder, even today, of the power of a strong personality and that the belief in “what they know to be true” can dramatically influence the direction of science and public opinion. I do not think that anyone would dispute that strong personalities and beliefs can influence science and public opinion. Take, for example, Julian Huxley’s rejection of Darwin’s ideas on sexual selection [11], undoubtedly reducing interest in that topic for the next 50 years or so.  And while it is true that Huxley and Cuvier had strong personalities, and were great communicators and relatively powerful men, I think that their arguments held sway largely because they made them clearly and because there was neither compelling evidence nor any clear and logical mechanisms to explain the existing patterns. In both cases the delays in the progress of science were reasonably short and probably needed the ideas and considerable evidence presented by Darwin and Wallace, and Williams and Trivers, respectively, before there could be any real progress.

Finally, it has probably not escaped your notice that the African Sacred Ibis has been depicted on the cover of The Ibis in one form or another ever since 1859. This may seem a bit odd as that bird does not occur in the wild in Britain and only sparsely in southern Europe through introductions. Thus the sacred ibis does not really appear to be a fitting symbol for the British Ornithologists’ Union. There is a long and interesting story there, but that too will have to wait for another day.

SOURCES

  • Birkhead TR, Wimpenny J, Montgomerie R (2014) Ten Thousand Birds: Ornithology since Darwin. Princeton, NJ: Princeton University Press.
  • Cuvier G (1804) Mémoire sur l’ibis des anciens Égyptiens. Annales du Muséum d’histoire naturelle 4:116-135. [available here]
  • Cuvier G (1812) Recherches sur les Ossemens Fossiles de Quadrupèdes : où l’on rétablit les caractères de plusieurs espèces d’animaux que les révolutions du globe paroissent avoir détruites, t1-4. [Studies of the Fossil Bones of Quadrupeds, volumes 1-4] Paris: Deterville. [available here]
  • Cuvier G (1826) Discours sur les révolutions de la surface du globe, et sur les changemens qu’elles ont produits dans le régne animal. Paris: G. Dufour. [available here and in English translation of the 1825 edition here]
  • Lacépède B-G-E, Cuvier G, Lamarck J-B (1802) Rapport des professeurs du Muséum sur les collections d’histoire naturelle rapportées d’Égypte, par E. Geoffrey. Annales du Museum d’Histoire Naturelle 1: 234–241. [available here]
  • Latham J (1790). Index Ornithologicus, Sive Systema Ornithologiae: Complectens Avium Divisionem In Classes, Ordines, Genera, Species, Ipsarumque Varietates (2 Volumes, in Latin). London: Leigh & Sotheby. [available here]
  • Le-Suer RB, ed (2012) Between Heaven and Earth: Birds in Ancient Egypt. Chicago: The Oriental Institute of the University of Chicago
  • Linnæus  C (1758) Systema naturæ per regna tria naturæ, secundum classes, ordines, genera, species, cum characteribus, differentiis, synonymis, locis, 10th edition.  Stockholm: Laurentius Salvius.
  • Rosser WH (1837) Mummy of Egyptian Ibis. The Gentleman’s Magazine 6 (new series): 145-148. [available here]
  • Savigny J-C (1805) Histoire naturelle et mythologique de l’ibis [Natural and Mythological History of the Ibis]. Paris: Allais. [available here]

Footnotes

  1. ibis described: see Latham 1790 page 706 where he calls it Tantalus aethyopius
  2. Sacred ibis once common in Egypt: but it is no longer found in that country, disappearing as the swamps and marshes were drained to provide land for the increasing population and agriculture.
  3. Late and Ptolomeic Periods: about 700 BC until 30 BC ending with the death of Cleopatra and the conquest of Egypt by the Romans
  4. ibises eating refuse: in Australia, where they have been introduced, they are often called ‘bin chickens’ as they are often seen foraging in trash cans in city parks. In the park beside the Australian National Museum in Sydney, I once watched a very dirty-looking ibis sneak up behind some picnickers then reach over the shoulder of a little boy to snatch his sandwich out of his hand. Clearly, their bills are adapted for sandwich snatching (!), and they are fearless.
  5. savants: these were scholars and scientists. The Journal des Sçavants (later called Journal des Savants) began publishing in January 1665, a couple of months before the Philosophical Transacations of the Royal Society, considered (erroneously) by many to be the first scientific journal. Frankly, I don’t see that it matters who was first, or even if it was one of those two.
  6. Grobert: (1757-181?) was a French artillery officer who wrote about the pyramids etc. on his return from the Egyptian campaign. See Cuvier (1826), which updates some of the information in his 1804 publication about his later study of some different mummies.
  7. smaller than contemporary ibises: On examining four more ibis mummies, Cuvier recognized that one of them was a juvenile based on its bone structure (Cuvier 1826). As a result, he realized that they may not be curlews at all but simply smaller, juvenile ibises. This is not so surprising as it turns out the many of the ibis mummies were clearly made from juvenile birds. No doubt the priest-farmers who raised the ibises for the votive market saw no reason to keep the birds any longer than was needed to make them suitable for mummification.  That just made good economic sense to maximize their profits.
  8. article in PLoS Biology: unlike the scientific articles in that journal, this one is labelled ‘Essay’ which they say “are opinionated articles on a topic of interest to scientists, as well as to a broader audience, including the general public”. Opinions are fine but I am surprised at the absence of clear evidence in support of the claims made.
  9. not actually a test: Cuvier simply used his measurements identifying the mummies as sacred ibises to suggest that there had not been much change in their morphology in the past 3000 years. But the ibis was just one of many examples that he referred to. Moreover, as Coyne noted, this was at best a ‘one-way test’ as any lack of change would be consistent with slow evolutionary change. Cuvier even acknowledged that the the measurements were not the same between the mummies and contemporary ibises. I don’t see this as a test of any kind because Cuvier was unlikely to be convinced by any such results: if the mummies were the same as extant ibises, then no change; if they were different then they must be different species.
  10. the story of Cuvier’s ibis measurements: in a quick search on the internet, I found more than 20 articles on Cuvier’s ibis measurements dating back to Rosser (1837)
  11. Huxley and sexual selection: see Birkhead et al. (2014)

NOTE As some of you may have noticed, this essay was briefly posted by accident in draft from a week ago. I immediately deleted that version from this blog and the final version above is substantially different, correcting several errors in the original and providing additional information, references and links.

Ladies, Parakeets, and the Biogeography of an Extinct Bird

BY: Bob Montgomerie, Queen’s University | 25 June 2018

In 1850, an anonymous author published a superb diary of natural history observations called ‘Rural Hours by a Lady’ based on two years of exploring the woods and fields near Cooperstown, New York. The book was wildly popular, and it was not long before the author was revealed to be Susan Fenimore Cooper [1]. On page 146 she says:

Parakeet2It is well known that we have in the southern parts of the country a member of the Parrot tribe, the Carolina Parakeet. It is a handsome bird, and interesting from being the only one of its family met with in a temperate climate of the Northern Hemisphere. They are found in great numbers as far north as Virginia, on the Atlantic coast; beyond the Alleghanies, they spread themselves much farther to the northward, being frequent on the banks of the Ohio, and in the neighborhood of St. Louis. They are even found along the Illinois, nearly as far north as the shores of Lake Michigan. They fly in flocks, noisy and restless, like all their brethren…In the Southern States their flesh is eaten…Birds are frequently carried about against their will by gales of wind; the Stormy Petrels, for instance, thoroughly aquatic as they are, have been found, occasionally, far inland. And in the same way we must account for the visit of the Parakeets to the worthy Knickerbockers about Albany.  [2]

Here, she correctly describes the bird as being most common in the southeastern states, though seen regularly as far north as the Great Lakes west of the Allegheny Mountains. What she did not know, of course, was that these were two subspecies, with different morphologies, ecologies and migratory strategies, as described below.

Live_captive_Carolina_parakeet
Shufeldt’s photo

The Carolina Parakeet was still abundant throughout its range in 1850 but, like the Passenger Pigeon, was soon to be extirpated. The second-last individual was a female called ‘Lady Jane’ who died at the Cincinnati Zoo in 1917; the last of its kind being Lady Jane’s mate, a male named ‘Incas’ who died there in 1918, one hundred years ago. Coincidentally, Incas died in the same cage where Martha, the last passenger pigeon, had died in 1914 [3]. There were reports of sightings in the wild for another 40 years or so, in Florida and Georgia, but none of those records were authenticated. Among the North American birds that have become extinct since the arrival of Europeans, the biology of the breeding biology Carolina Parakeet may be the poorest known [4]. And there is, surprisingly, only one photo of the bird in a natural-looking setting [5], taken by the irrepressible Robert Shufeldt in about 1900.

 

A recent pair of papers by Kevin Burgio and colleagues uses all of the known specimens and sightings of this bird to reveal some interesting insights into its distribution, ecology, and taxonomy. There were 401 of those sightings recorded between 1564 and 1944, and nearly 800 specimens in museums and private collections worldwide [6], almost all collected in the 1800s. As shown on the graphs below, the number of records climbed exponentially from 1500 to 1900, reflecting the increases in exploring the new continent, in writing about natural history, and in preserving ornithological data and specimens. There was an uptick in collecting, or at least preserving specimens, from 1870-1900 when it became clear that the bird was disappearing [6].

graphs
Records and specimens with known dates—note the log scale on upper two graphs.

Analyzing records only from states where the parakeet was known to breed, Burgio and colleagues, georeferenced all the data and used 147 unique localities to create the species breeding distribution models shown on the map to the right below. The map on the left was produced in 1889 by Edwin Hasbrouck with the known range in his day (black shading) nicely matching the newly reconstructed ranges of the two subspecies.

Burgio and colleagues’ research also suggested (i) that the breeding range of this species was much smaller than previously thought, (ii) that the two subspecies, previously only vaguely defined by size and colour, actually had disjunct ranges and occupied somewhat different climatic niches, and (iii) that the western subspecies was almost certainly migratory where the eastern one was not.

The authors also hoped their analysis would help to inform current conservation practices in an effort to save the 8% of bird species currently threatened to disappear as a result of climate change. Parrots, in particular, are in bad shape, with 42% of species listed as threatened or endangered.

MAPS
LEFT from Hasbrouck (1889) estimating the limits of the parakeet’s historical range (black line) with shading showing the range in the 1880s. RIGHT from Burgio et al. (2017) estimating the ancestral breeding ranges of the two subspecies (Hasbrouck’s range limit shown as a red line)

 

pipe
Clay pipe ca. 1650

As much as I like those recent papers, I think it’s unfortunate that many biogeographers draw their maps as if animals obeyed political boundaries, as on the right-hand but not the left-hand maps above. The right-hand graph implies, for example, that the bird could never have crossed the US-Canada border as there was nowhere to go. Despite that, there is some evidence that it did occasionally occur in southwestern Ontario, possibly blown off course as Susan Cooper suggested above. At an archaeological dig at Grimsby, Ontario, for example, Walter Kenyon found a clay pipe that looks distinctly like a parrot, made by native peoples in the mid-1600s. And Rosemary Prevec found 3 Carolina Parakeet bones at a native site near London, Ontario, dated at around 1100 CE. Both of these findings are no more than suggestive and could have been obtained in trade with natives living further south.

Possibly more convincing are some observations that Samuel de Champlain recorded in his notes in 1615, in the woods near where I live in Kingston. He says that he  “…penetrated so far into the woods in pursuit of a certain bird which seemed to be peculiar, with a beak almost like that of a parrot, as big as a hen, yellow all over, except for its red head and blue wings, which made successive flights like a partridge.” [8] There are definitely no other birds even remotely resembling that description in eastern Ontario today.

fancyfeathers
Catalog page, New York Millinery and Supply Company, Inc., New York

None of this nationalism is really important to our understanding of the bird’s ecology and demise, except to note that at one time the species was clearly widespread and mobile. What is important is an attempt to understand why they went extinct, as even by the middle of the 1800s it appeared to be declining in numbers [6].

Burgio and colleagues point to habitat destruction and hunting as the likely causes. Not surprisingly, the parakeet’s feathers were prized for the millinery trade, with some reports suggesting that ladies hats were sometimes adorned with skins of the whole bird. The 1901 ad to the right, for example, shows a whole parrot (skin) in the lower right corner, for the bargain price of 25¢ a bird or $2.95 a dozen (about $7.50 and $88 in today’s currency). While the documentation is sketchy, it is also likely that this species was a popular cage bird in Germany as well as in North America. The only other known photo, besides Shufeltdt’s, is also one of a pet called  ‘Doodles‘, kept by Smithsonian malacologist Paul Bartsch. In 1900, ‘doodle‘ meant ‘fool‘ and not the ‘absentminded scribble‘, Google commemorative, or online scheduler that it is today. And I wonder if Bratsch gave it that name to reminder him what fools we are when let any species go extinct.

SOURCES

  • Anonymous [Cooper, SF] (1850) Rural Hours by a Lady. New York: G. Putnam.
  • Burgio KR, Carlson CJ, Tingley MW (2017) Lazarus ecology: Recovering the distribution and migratory patterns of the extinct Carolina parakeet. Ecology and Evolution 7:5467–5475.
  • Burgio K, Carlson C, Bond A (2018) Georeferenced sighting and specimen occurrence data of the extinct Carolina Parakeet (Conuropsis carolinensis) from 1564-1944. Biodiversity Data Journal 6:e25280.
  • Cokinos C (2000) Hope Is the Thing With Feathers: A Personal Chronicle of Vanished Birds. New York: Penguin.
  • Fuller E (2013) Lost Animals: Extinction and the Photographic Record. London: Bloomsbury.

  • Greene WT, Dutton FGFG, Fawcett B, Lydon AF (1883) Parrots in Captivity, v. 2. London: George Bell and Sons.

  • Hahn P (1963) Where is that Vanished Bird? Toronto: University of Toronto Press. [see this previous post for more on this book[
  • Kennedy CC (1984) Did Champlain stalk a Carolina Parakeet in southern Ontario in 1615? Arch Notes 84:55–62.

  • McKinley, D. (1960) The Carolina parakeet in pioneer Missouri. The Wilson Bulletin 72:274–287.
  • McKinley D (1977) Climatic relations, seasonal mobility, and hibernation in the Carolina Parakeet. Jack-Pine Warbler 55:107–124.
  • Prevec R (1984) The Carolina Parakeet—its first appearance in southern Ontario. Arch Notes 84:51-54.
  • Snyder NFR (2004) The Carolina Parakeet: Glimpses of a Vanished Bird. Princeton, NJ: Princeton University Press.

Footnotes

  1. Susan Fenimore Cooper: was a superb naturalist, author, and artist whose work was overshadowed in more ways than one by that of her father, James. She deserves recognition and a separate essay on her own work, Stay tuned.
  2. Cooper quotation: from Anonymous 1850 page 146
  3. ‘Incas’ the parakeet: Like Martha, the last Passenger Pigeon, Incas was frozen and sent to the Smithsonian, but he was lost in transit (Fuller 2013)
  4. breeding biology poorly known: see Snyder (2004)
  5. Shufeldt’s photo: is one of a pair of pet birds that Shufeldt borrowed from his friend Edward Schmidt, and it took him hours to get it to sit still enough on a cocklebur to make a decent photo (Cokinos 2001). Both of Schmidt’s birds later died from chewing on the bars of their cage, possibly from lead paint poisoning (Fuller 2013)
  6. declining numbers by mid 1800s: see Hasbrouck (1889)
  7. records and specimens: see Hahn (1963), McKinley (1960, 1977) and Snyder (2004) for background
  8. Champlain quotation: from Kennedy 1984 page 55

IMAGES:  first parakeet is by Robert Ridgway from Baird et al. (1905); graphs by the author based on data in Burgio et al. (2017, supplement)—parakeet is an engraving by Benjamin Fawcett in Greene et al. (1883); maps from the original papers; clay pipe from Kennedy (1984); millinery ad from the Smithsonian collection

Why Woodpeckers are Scarce in the North

BY: Bob Montgomerie, Queen’s University | 18 June 2018

On the 18th of June 1858, one hundred and sixty years ago today, Darwin claims [1] to have received that fateful letter from Alfred Russel Wallace—probably the most famous letter in the history of science. The original letter was lost but it was transcribed and read to the Linnean Society of London on 1 July and published later in their journal. That letter is well worth reading, especially because it contains some interesting insights into avian ecology. While Wallace had some useful ideas relevant to natural selection, it could be argued that his argument was not nearly as well-formed as Darwin’s [2]. In a way, his ecological and biogeographical insights are more original, in my opinion.

Alfred_Russel_Wallace_1862_-_Project_Gutenberg_eText_15997
Wallace in 1862

Wallace wrote that letter on Ternate in the Mollucas in February 1858, sent it out on a mail steamer on 5 April. He was in the South Pacific for 8 years on a collecting trip, in part to obtain specimens that he could sell back in England but also to gather material for books that he thought, rightly so, would provide him with a lifetime of fame and fortune. He brought home more than 125,000 specimens, including more than 8000 bird skins.

 

I found three things to be remarkable about Wallace’s letter. First, he develops some of the same ideas about selection as Darwin,  and uses some of the same language: “state of nature”, “struggle for existence”, “stability of species”, “geometrical ratio”, “origin of…species”,  “conditions of existence”, and “superior varieties.” These are not phrases you would be likely to read in a recent paper on evolutionary biology, but may well have been argots of the scientific literature in the 1800s.

Second, he makes clear his objections to Lamarck’s ideas:

The hypothesis of Lamarck—that progressive changes in species have been produced by the attempts of animals to increase the development of their own organs, and thus modify their structure and habits—has been repeatedly and easily refuted…the view here developed renders such an hypothesis quite unnecessary, by showing that similar results must be produced by the action of principles constantly at work in nature. [3]

And third, he is remarkably insightful and creative about what today we would call evolutionary ecology with respect to passenger pigeons, woodpeckers, and clutch size.

On clutch size, he makes the perceptive observation that a species’ population size—and rate of increase—has nothing to do with the number of offspring in a brood:

…large broods are superfluous. On the average all above one become food for hawks and kites, wild cats and weasels, or perish of cold and hunger as winter comes on. This is strikingly proved by the case of particular species; for we find that their abundance in individuals bears no relation whatever to their fertility in producing offspring. Perhaps the most remarkable instance of an immense bird population is that of the passenger pigeon of the United States, which lays only one, or at most two eggs, and is said to rear generally but one young one. [3]

GouldGSWoopecker
Great Spotted Woodpecker

On woodpeckers, he argues that they are more scarce in the temperate zone than in the tropics due to the uncertainty of overwinter food in the north, and the various morphological adaptations that would make long-distance migration difficult. I don’t even know if these observations are true, but the idea is immensely creative and demonstrates his excellent ecological insights:

 

Those whose organization does not permit them to migrate when their food becomes periodically scarce, can never attain a large population. This is probably the reason why woodpeckers are scarce with us, while in the tropics they are among the most abundant of solitary birds. Thus the house sparrow is more abundant than the redbreast, because its food is more constant and plentiful,- seeds of grasses being preserved during the winter, and our farm-yards and stubble-fields furnishing an almost inexhaustible supply. [3]

On the Passenger Pigeon, he reasons—correctly, I think—that its unbelievably huge populations were a product of the bird’s ability to move efficiently to track the vagaries of its occasionally superabundant food supply:

Perhaps the most remarkable instance of an immense bird population is that of the passenger pigeon of the United States, which lays only one, or at most two eggs, and is said to rear generally but one young one. Why is this bird so extraordinarily abundant, while others producing two or three times as many young are much less plentiful? The explanation is not difficult. The food most congenial to this species, and on which it thrives best, is abundantly distributed over a very extensive region, offering such difference of soil and climate, that in one part or another of the area the supply never fails. The bird is capable of a very rapid and long-continued flight, so that it can pass without fatigue over the whole of the district it inhabits, and as soon as the supply of food begins to fail in one place is able to discover a fresh feeding-ground. [3]

Like his contemporaries, however, Wallace reasoned that this species’ populations were just too big to fail: “This example strikingly shows us that the procuring a constant supply of wholesome food is almost the sole condition requisite for ensuring the rapid increase of a given species, since neither the limited fecundity, nor the unrestrained attacks of birds of prey and of man are here sufficient to check it. In no other birds are these peculiar circumstances so strikingly combined.” [3] This is one of those rare cases where we could actually learn from history and maybe not repeat Wallace’s mistake in our dealings with other species.

SOURCES

  • Bock WJ (2009) The Darwin-Wallace myth of 1858. Proceedings of the Zoological Society 62:1–12.

  • Davies R (2008) The Darwin conspiracy: origins of a scientific crime. London: Golden Square Books
  • Davies R (2012) How Charles Darwin received Wallace’s Ternate paper 15 days earlier than he claimed: a comment on van Wyhe and Rookmaaker (2012). Biological Journal of the Linnean Society 105:472–477.

  • Darwin CR, Wallace AR (1858) On the tendency of species to form varieties; and on the perpetuation of varieties and species by natural means of selection. Journal of the Proceedings of the Linnean Society of London, Zoology 3:46–50.

  • Davies R (2012) How Charles Darwin received Wallace’s Ternate paper 15 days earlier than he claimed: a comment on van Wyhe and Rookmaaker (2012). Biological Journal of the Linnean Society 105:472–477.

  • Gould E, Gould J, Lear E (1837) The Birds of Europe. (v. 1-5). London: pub. by the author.
  • Smith CH (2013) A further look at the 1858 Wallace–Darwin mail delivery question. Biological Journal of the Linnean Society 108:715–718.


Footnotes

  1.  Darwin’s claim about Wallace’s letter: Davies (2008) in particular, claimed that Darwin received the letter earlier and plagiarized it in his own notes so that he could claim priority, This seems highly unlikely to me, based on what I know of Darwin’s character and what Darwin himself says about the letter. van why and Rookmaaker (2012) present a convincing counter argument (but also see Davies 2012)
  2. Wallace’s ideas on natural selection: see Bock (2008) for details on what Wallace did have to say about selection
  3. Quotations: are all from the transcribed version of Wallace’s letter, available here

The Spring Rivalry of Birds

BY: Bob Montgomerie, Queen’s University | 28 May 2018

Unless you have been living in a cave for the past couple of months, you will have been well aware of the spectacle of mature, powerful males posturing to one another, showing off their weapons, advertising their prowess, and intimidating their neighbours. Some of them even coerced females into unwanted sexual acts, or spent some time cheating on their mates. I refer, of course, to what Charles Bethune Moffat called ‘The Spring Rivalry of Birds‘ in his 1903 paper in The Irish Naturalist.

CootFight
American Coots fighting at a territory boundary (PHOTO Bruce Lyon)

Moffat was born in January 1859 on the Isle of Man in the Irish Sea, 10 months before the publication of Darwin’s Origin of Species. Moffat’s family soon moved to Ireland where he grew up and eventually spent his entire professional career as a lawyer and journalist.

IrishNatBut Moffat was also a consummate naturalist who published about birds in popular articles in Dublin’s Daily Express, and in scientific papers in The Irish Naturalist [1] and The Irish Naturalists’s Journal. His paper on the ‘spring rivalry’ is best known because it was probably the first publication to present, in English [2], the idea that male birds defend a territory: “…that the battles fought between the male birds in spring have for their object, not the winning of particular females, but the acquiring of suitable plots of land, and that the song and bright plumage of the male are a warning to other males and an intimation to a female that a suitable territory has been acquired.” [3]

Reading Moffat’s spring rivalry paper, I get the distinct impression of a man who was thoughtful, logical and a keen observer—all traits that would have made him an excellent lawyer and journalist (both of which he was) and a really good scientist. He noted, for example, “…that we seldom find in close proximity to each other two nests belonging to the same species of bird.”, “…that cock birds in early spring spend a great deal of their time in fighting one another.”, and “..that we must have a very large number of non-breeding birds of both sexes, prevented from breeding simply by the fact that they have no suitable ground.” [3]

Moffat liked Darwin’s ideas about evolution and natural selection but he thought—like many others at the time—that Darwin was simply wrong about sexual selection, especially with regard to female choice:

…the number of hen-birds, so far as we can gather, is fully equal to the number of cock-birds, so that when all the fighting is over, there is nothing to prevent all the birds from marrying and settling down to ‘ live happily ever after.’ We may suppose, in our sentimental way of looking at things, that even then the poor beaten cock-bird suffers from a certain amount of depression when he thinks of the greater charms of her to whom he first paid court. But we have the assurance of experts that no such thing happens; that one hen-bird is quite as good as another, and that every cock-bird is perfectly content with the first mate he can get. That is sometimes laid down as the reason why hen-birds, as a rule, have not developed bright nuptial colours or melodious voice…I think we must infer that there is very little free choice or aesthetic selection, and that the hen bird is mainly guided by prudential motives in accepting the owner of the soil. [3]

JuncoFight
Dark-eyed Junco fighting his reflection in a car window (PHOTO: Bruce Lyon)

Having observed male Blackbirds and Chaffinches continually battling their reflections in his windows, Moffat quite perceptively concluded that these fights were about space, as there was no female in sight and those reflected interlopers had nether sung nor shown any interest in the male’s mate: “the imaginary enemies, on whom so much fury was expended, were guilty of no crime beyond that of being in the spot where they were.” [3]. Even when 35 Starlings were shot one at a time, consecutively, from a single breeding pair, the remaining bird almost immediately got a new partner, suggesting to Moffat that birds did not really care who they mated with, and that space was therefore of primary importance.

But why ‘space’? Moffat was also skeptical about Darwin’s idea that—at least in birds—many more offspring are produced than could possibly survive to reproduce: “I cannot believe that the theory of Natural Selection — for which I have a great respect, and which I must carefully guard myself against appearing for a moment to call in question— requires this sacrifice, or anything like it…as regards birds I am altogether unable to find grounds for believing in so great a death-rate..” [3]. Indeed, his censuses of his local House Martins suggested to him that the annual mortality rate was very low, for just about as many birds arrived back in spring as had departed the previous autumn [4].

VCWE1962His solution to this conundrum of territoriality—anticipating V. C. Wynne-Edwards’s ideas on group selection published 60 years later—was: “..that there are checks of a prudential kind on the marriage of birds, and that these checks may be a very important factor in keeping the number of birds absolutely permanent.” [3]. Thus Moffat argued that birds were simply being prudent, dividing up the available land in such a way that would keep the population stable, without the needless waste of reproduction wherein the excess individuals would die off as Darwin had argued.

He was certainly not alone in his skepticism about sexual selection and population regulation, and it took the reaction to Wynne-Edwards’s ideas—mainly by David Lack and George Williams—to set the record straight with regard to the logic of natural selection. And that set the stage for a re-examination of sexual selection and its influence on the songs, plumage colours and ornaments, and courtship behaviours of birds—traits that benefit individuals not species.

So, next time you hear the twitter of North America’s Orange-crowned Warbler, see European Coots threaten their neighbours, watch the Siberian Grouse perform his flutter-jump, or hear the Oriental Cuckoo whistle his ridiculous ‘poo-poo’, remember that none of this male behaviour is performed for the good of the species.

SOURCES

  • Altum B (1868) Der Vogel und sein Leben. Münster: Niemann. [available here]

  • Kennedy PG (1946) Charles Bethune Moffat. (1859-1945). British Birds 39:81–82.

  • Moffat CB (1903) The spring rivalry of birds. Some views on the limit to multiplication. The Irish Naturalist 12:152–166.

  • Stresemann E (1947) Baron von Pernau, pioneer student of bird behavior. The Auk 64:35–52.

  • von Pernauer FJA [published anonymously without his consent] (1702) Unterricht, was mit dem lieblichen Geschöpff, denen Vögeln, auch ausser dem Fang, nur durch die Ergründung deren Eigenschafften und Zahmmachung oder anderer Abrichtung, man sich vor Lust und Zeitvertreib machen könne, gestellt Durch den Hoch- und Wohlgeborhrnen Hn. Freyherrnn: Herrn von P [ available here]
  • White G (1772) Letter XI to The Honourable Daines Barrington. [available here]

  • Wynne-Edwards VC (1962) Animal Dispersion in Relation to Social Behaviour. Edinburgh: Oliver and Boyd.


Footnotes

  1. papers in Irish Naturalist: he published 40 papers and short notes in this journal, as well as 40 book reviews
  2. first presentation in English: Ferdinand Johann Adam von Pernauer (1702) and, especially, Bernard Altum (1868) had previously introduced and developed the idea of territoriality and the concept of ‘territory’ in birds. Moffat also notes that Gilbert White had a similar view that he expressed in a letter written in 1772: “such a jealousy prevails between the male birds that they can hardly bear to be together in the same hedge or field…it is to thisspirit of jealousy that I chiefly attribute the equal dispersion of birds in the spring over the face of the country.”
  3. quotations: all from Moffat (1903)
  4. House Martin observations: his mistake here is that he assumed that these were the same individuals, not realizing that overwinter mortality would have left openings to be filled by birds that might otherwise have been non-breeders

Lumpers and Splitters

BY: Bob Montgomerie, Queen’s University | 14 May 2018

Almost everyone I know who is interested in birds has their favourite group, and the reasons for those favourites vary widely. For some, the difficulty—and their mastery—of field identification is most appealing: I am thinking peeps, LBJ sparrows, fall wood warblers in North America and leaf (Phyloscopus) warblers in Europe. For others, it’s the beauty of—variously—their songs (wood warblers, thrushes, mimics), their plumages (male wood warblers in spring, hummingbirds, trogons, peafowl), their ability to fly (falcons, hummingbirds, swallows, swifts), their intelligence (corvids, tits, parrots), or their way of life (seabirds, hummingbirds, waterfowl, shorebirds, antbirds, brood parasites, cooperative breeders, lekking species, owls).

peeps
Peeps at Moss Landing, California

For many scientists, their favourites are the birds that they have studied most—often as much by accident as by design. Among my ornithologist friends and colleagues, the favourites are coots, murres, manakins, Parids, fairy wrens, acorn woodpeckers, barn swallows, anis, and snow geese. My own favourites are hummingbirds (PhD research), shorebirds and ptarmigan (20 years of arctic research), and seabirds (summer job before PhD).

When I began birding, however, my favourites were the Hylocichla (now Catharus) thrushes and the Empidonax flycatchers, possibly because some species were hard to identify. I was also intrigued by the geographic mosaic of Empidonax ranges and the incredible similarities in their plumages described by Ned Johnson, among others. Why is it that some species in some genera (Dendroica males in springfor example) are so easy to tell apart whereas in other genera (like Empidonax) species can only be distinguished morphologically by careful measurements of a bird in the hand (or so we thought). My own careful examination of Empidonax flycatchers while banding them at Long Point Bird Observatory resulted my first paper in an international journal more than 50 years ago.

WillowL_Alder_060107a-1tbh9a8
Willow (L) and Alder (R) Flycatchers

Because of that early interest in the Empidonax flycatchers, I was both dismayed and intrigued when the AOU split the Traill’s Flycatcher into two species—Alder and Willow—in 1973 on the basis of their song and some morphological traits. Here was a species that I thought I knew well (albeit only on migration), that was actually two species, even more cryptic than the other hard-to-tell-apart species in that genus. This was my first lesson in the fluidity of taxonomy and the possibility that there were almost certainly other cryptic species to be found. I remember wondering, for example, if the Grey-cheeked Thrush might be two species, based on the obvious morphological and geographic differences in the subspecies bicknelli, and that, too, was formally recognized as a separate species by the AOU in 1995.

Throughout the 20th century, there was often a tension between lumpers and splitters [1] when it came to avian taxonomy. Some suggested that the differences between splitting and lumping were like the differences between liberal and conservative governments in democracies. We now recognize, however, that periods of lumping and splitting are the result of changing opinions about species definitions, and the progress of technologies that allow us to distinguish among populations.

A new paper in PLoS ONE by Gaurav Vaidya, Denis Lepage, and Robert Guralnick [2] uses the 136 years of AOU checklists to examine, among other things, the historical patterns of lumping and splitting of the North American avifauna by taxonomists. The first AOU checklist was published in 1886, the production of which was one of the stated reasons for the for the formation of the American Ornithologists’ Union in 1883. Since then, 6 additional full ‘editions’ of the checklist have been published, with 57 supplements published at roughly 2-year intervals between editions.

Since 1886, fully three-quarters of the ~900 North American species have not been involved in what the authors call the ‘correction process’, being either lumped with other species or split into two or more species by the checklist committees. Sixteen percent have been corrected once, and the remaining 10% twice or more. In all, there have been more  lumps (142) than splits (95) in the past 130 years but the pattern of change has been striking. Lumping was initially the norm, probably largely due to the widespread adoption of the Biological Species Concept. But very few species have been lumped since 1980 as the rate of splitting took off, likely because DNA technologies improved and there began a (slight) shift toward a phylogenetic species concept.

VaidyaGraph

Their analysis identified three periods of relative stasis, when there were no corrections to be made, perhaps roughly coinciding with periods of societal upheaval and dramatic scientific changes in taxonomic philosophies and practices.

Vaiyda and colleagues also identify some intriguing examples of  “… a current, ongoing taxonomic recorrection process, in which corrections made in the first half of the 20th century are now being reverted in light of new evidence and better tools. [3]. To the non-scientist this must look like the taxonomists are incredibly indecisive but these recorrections actually (often) reflect the sort of changes in species definitions and technologies described above. For example, in 1923 the Common Galllinule (Gallinula galeata) of North America was lumped with the Common Moorhen (Gallinula chloropus) of Europe because they were thought, based on their breeding ranges and morphologies, to be subspecies. Then, in 2011, those species were split again, recognizing the differences in their vocalizations, bill and shield morphologies, and mtDNA.

GallinulaCompare
Common Gallinule (L) and Common Moorhen (R)

If the trend shown in the graph continues—and there is no reason to expect that it will not—the number of splits will soon surpass the number of lumps. Recently, George Barrowclough and colleagues have speculated that the level of splitting resulting from the application of genomic tools and the phylogenetic species concept will almost double the number of species recognized worldwide—to 18,000—before too long. The implications of this for conservation, for understanding the evolutionary history of birds, and for my life list, are huge.

SOURCES

  • American Ornithologists’ Union. 1995. Fortieth supplement to the American Ornithologists’ Union Check-list of North American Birds. Auk 112: 819-830.

  • Barrowclough GF, Cracraft J, Klicka J, Zink RM (2016) How many kinds of birds are there and why does it matter? PLoS One. 2016; 11: e0166307. https://doi.org/10.1371/journal.pone.0166307 PMID: 27880775
  • Chesser RT, Banks RC, Barker FC, Cicero C, Dunn JL, Kratter AW, Lovette IJ, Rasmussen PC, Remsen Jr JV, Rising JD, Stotz DF, Winker K. (2011) Fifty-Second Supplement to the American Ornithologists’ Union Check-List of North American Birds. Auk 128: 600-613
  • Johnson NK (1980) Character variation and evolution of sibling species in the Empidonax difficilis-flavescens complex (Aves: Tyrannidae). University of California Publications in Zoology 112: 1-151.

  • Mayr E (1982) The Growth of Biological Thought. Cambridge, Mass: Belknap Press.

  • Stone W, Oberholser HC, Dwight J, Palmer TS, Richmond CW (1923) Eighteenth Supplement to the American Ornithologists’ Union Check-List of North American Birds. Auk 40:513–525.

  •  Vaidya G, Lepage D, Guralnik R (2018) The tempo and mode of the taxonomic correction process: How taxonomists have corrected and recorrected North American bird species over the last 127 years. PLoS ONE 13(4): e0195736. https:// doi.org/10.1371/journal.pone.0195736


Footnotes

  1. lumpers and splitters: respectively, taxonomists who favour placing similar populations under the umbrella of a single species versus those who seek to identify distinct populations as separate species. Ernst Mayr (1982, page 240) claimed that “As a general rule one can say that most taxonomic groups pass through a phase of rather intensive splitting when they are studied more actively, but that the splitting phase is reversed when the knowledge of the group reaches greater maturity.” I think this new paper by Vaidya et al. (2018) shows that statement to be incorrect and may reflect Mayr’s view that his Biological Species Concept was sacrosanct.
  2. while this is an excellent analysis the paper is technically detailed, a necessity given the changes in the way species have been defined and the changes in coverage of the checklist. I have tried to distill some of those details here but I am no taxonomist.
  3. quotation: from Vaidya et al (2018 page 2)

IMAGES: peeps by the author; graph redrawn by the author from Vaidya et al (2018, Figure 1); Flycatchers from https://u.osu.edu/biomuseum/2015/09/28/examples-of-sibling-species/comment-page-1/; Gallinula spp. from Wikimedia Commons, in the public domain.

Professor Bumpus and his Sparrows

Guest Post

BY: Ted R. Anderson | 5 March 2018

Possibly the most influential ornithological paper published inNorth America in the 19th century was actually written by an invertebrate embryologist who was not even a member of the American Ornithologists’ Union.  The paper “The elimination of the unfit as illustrated by the introduced sparrow, Passer domesticus” was written by Professor Hermon Carey Bumpus at Brown University in Providence, Rhode Island  It was actually the second of two interesting papers that Bumpus published on the recently introduced house sparrow, but more on these below.

bumpus
Hermon Bumpus

Bumpus was born in Maine in 1862, and entered Brown in 1879 to study biology, graduating in 1884.  In 1886, he accepted a professorship at Olivet College in Michigan, a position he left in 1889 to complete a doctorate at the newly established Clark University, where he received the first PhD awarded by that university.  In 1890 he returned to Brown as assistant professor of zoology and was promoted to professor of comparative anatomy two years later.  He left Brown in 1900 to become assistant to Morris Jessup, president of the board of the American Museum of Natural History in New York City.  A year later Jessup promoted him to become the museum’s first director.

In 1911 Bumpus moved into academic administration as business manager of the University of Wisconsin, a position he held until 1914.  He then moved to Tufts College (now University) as President from 1915 to 1919.  He resigned from Tufts to pursue his interest in building or remodeling homes including a Philippine bungalow on Long Island Sound (constructed of Philippine lumber from the Philippine Hall at the St. Louis Exposition), an Italian villa in a Boston suburb and the King Caesar House in Duxbury, Mass.  Bumpus died in Pasadena, California in 1943.

pexels-photo-460960.jpegWhile teaching at Brown, Bumpus spent his summers conducting research on the development of marine invertebrates at the Marine Biological Laboratory at Woods Hole, Massachusetts, where he also served as assistant director from 1893 to 1895, and as director of the Biological Laboratory of the US Fish Commission.  In 1896 he presented the inaugural lecture in a summer seminar series at the Marine Biological Laboratory entitled “The variations and mutations of the introduced sparrow, Passer domesticus”, which was published in 1898.

In that 1898 paper he was undoubtedly the first scientist to suggest that the introduction of house sparrows and their subsequent rapid spread across North America represented a huge experiment that could be used to study Darwinian natural selection.  Taking advantage of that natural experiment, he compared the size, shape and coloration of 868 sparrow eggs from Massachusetts with an equal number of sparrow eggs from England, to test the hypothesis that the rapid population growth of sparrows in North America would result in relaxed selection.  Without the benefit of statistical analysis—Francis Galton and Karl Pearson were just then developing some rudimentary statistical tests—he concluded from his graphs that eggs from Massachusetts were shorter and more variable in size and coloration than eggs from England.  He also raised the question of whether the observed differences were phenotypic (‘ontogenetic’) or adaptive (‘phylogenetic’) and suggested that a common garden experiment would be needed to differentiate between these alternatives.

BumpusEggLength
Bumpus’s graph of the length of house sparrow eggs from North America (dotted line) and Europe (solid line) [1]

On 1 February 1898, a winter storm in Providence provided Bumpus with the material for another summer lecture at Woods Hole, which he then published.  After the storm, 136 immobilized sparrows were brought to Bumpus’s anatomy lab, where 72 subsequently revived but the remaining 64 died.  Bumpus identified the sex and measured nine morphological traits of each bird.  Bumpus concluded from his graphs that males survived better than females and that shorter, lighter birds with longer legs, wings and sternums and larger brain size (“skull width”) also survived better. He concluded that his analyses showed:

Natural selection is most destructive of those birds which have departed most from the ideal type, and its activity raises the general standard of excellence by favoring those birds which approach the structural ideal.

…the birds which perished have certain average structural peculiarities which distinguish them from the survivors, and that the intensity of selective elimination has been felt most by birds of extreme structure [2]

In his 1899 publication, the entire dataset is reproduced in an appendix, thereby permitting many other evolutionary biologists, as well as innumerable students in evolution classes, to analyze Bumpus’s data statistically.  Harris published the first  statistical analysis, and at least ten other papers have been published since then, including papers by John Calhoun, Peter Grant , Richard F. Johnston and colleagues, and one by Russell Lande and Steven Arnold.  Increasing complex and sophisticated statistical analyses were employed in these papers, and the conclusions of the various authors differ from those of Bumpus and from each other, in part due to the fact that many of the analyses use only subsets of the original data.

I do not know of another dataset of birds that has been subjected to so many analyses and so many different interpretations The history of reanalysis of Bumpus’s data is a nice example of a century of progress in both statistics and evolutionary biology

SOURCES

  • Anderson TR (2006) Biology of the Ubiquitous House Sparrow, from Genes to Populations. Oxford: Oxford University Press
  • Bumpus HC (1898) The variations and mutations of the introduced sparrow, Passer domesticus. Biological Lectures Delivered at the Marine Biological Laboratory of Woods Holl, 1896-1897, pp. 1-15.
  • Bumpus HC (1899) The elimination of the unfit as illustrated by the introduced sparrow, Passer domesticus. Biological Lectures from the Marine Biological Laboratory of Woods Holl, Mass. 1898, pp 209-228.
  • Calhoun JB (1947) The role of temperature and natural selection in relation to the variations in size of the English sparrow in the United States. American Naturalist 81:203-228.
  • Grant PR (1972) Centripetal selection and the house sparrow. Systematic Zoology 21:23-30.
  • Harris JA (1911) A neglected paper on natural selection in the English sparrow. American Naturalist 45:314-319.
  • Johnston RF, Niles DM, Rohwer SA (1972) Hermon Bumpus and natural selection in the house sparrow Passer domesticus. Evolution 26:20-31.
  • Lande R, Arnold SJ (1983) The measurement of selection on correlated characters. Evolution 37:1210-1226.

Footnotes

  1. graph: Bumpus 1898 page 5
  2. quotation: Bumpus 1899 pages 217 and 218

2017: An historic year for ornithology

BY: Bob Montgomerie, Queen’s University | 1 January 2018

The year that ended yesterday was an historic one for ornithology, with the first meeting of the newly formed American Ornithological Society (AOS) in East Lansing, Michigan. As a student of the history of ornithology, I know how hard it is to predict the future of our discipline (i.e. impossible), or to even guess correctly how well research published during the past year will be regarded 10, 50 or 100 years from now. In 1981, Bill Gates is alleged to have said that he did not see why anyone would ever need more than 640 kb of memory in a personal computer. While that story may be apocryphal, it undoubtedly resonates because it reflects our personal experience with crystal ball gazing.

Even so, it is hard for me to imagine that the new AOS is not a move in the right direction, maybe eventually embracing ornithological societies throughout the western hemisphere. We have already reaped some of the benefits of that merger in the new journal structure, new websites and social media, and new initiatives for funding and outreach. More on the 2017 AOS conference below, but first a very personal listing of a few events in 2017 that made news in ornithology.

People

The history of ornithology is, in my opinion, best viewed through the lives of people who study birds, as it is they who made the discoveries and often inadvertently set a course for the future. The Auk and The Ibis, as well as other journals, regularly publish formal obituaries of ornithologists, but I am hoping in the future that we can publish some more personal accounts on this blog. Sometimes—but too rarely—personal accounts do get published in our journals (see for example, Jon Ahlquist’s obituary for Charles Sibley).

I am also hoping that many senior ornithologists will contribute their own memoirs to a series we will be launching this year at the AOS meeting in Tucson. I did not know personally all of the ornithologists who died in 2017, but each of the following influenced me and my research in profound ways: Amotz Zahavi (b 1928), Patrick Bateson (b 1938), Wesley (Bud) Lanyon (1926), Harry Carter (b 1956), François Vuilleumier (b 1938), and Chandler Robbins (b 1918). All of them have or will likely soon be remembered in more formal obituaries in the journals—I have listed a few under Sources, below—but I would be happy to publish more personal accounts on this blog. Send me an email if you are interested in contributing.

Publications

ScienceCover2017Birds were on the covers of the highest profile journals of general science (Science, PNAS, but—notably—not Nature) and biology (Current Biology, Proceedings of the Royal Society B) no less than 8 times in 2017, reflecting the continuing interest and influence of ornithology.

I estimate that there were about 21,000 papers published on birds in 2017. That’s more papers than were published on birds from Aristotle to the beginning of the 20th century. Clearly nobody could (or would want to) keep abreast of the ornithological literature the way that Elliott Coues, Alfred Newton and Casey Wood once did in the late 1800s and early 1900s. That makes me wonder if scientists now rely more than ever on books and review articles to get a sense of their discipline.

AOS 2017

At the inaugural meeting of the AOS, in early August 2017, the slide show below was used to introduce the society and conference. Jen Owen (Michigan State University) put this show together to show some of the early history of the AOU and COS, and their development that culminated in the modern AOS.

My first AOU meeting was in Duluth, Minnesota, in 1966 when I was still a teenager. I travelled from Toronto to Duluth with my Royal Ontario Museum friends and mentors, Jim Baillie and Rolph Davis, in Rolph’s brand new Ford Mustang, getting stopped only once for speeding! While I felt very welcome at that conference, and learned a lot, I was taken aback that there were so few young people there and so few women (see also here). That was the year that the AOU first offered their Marcy Brady Tucker Award to provide travel subsidies ‘to assist a few promising young ornithologists’ to attend the annual meeting. I am not sure how many awards were given in 1966 (I did not apply) but I feel that was the beginning of a new era for the AOU.

I have not been to all of the AOU meetings in the intervening 50 years, but the 2017 AOS meeting was certainly one of the best of a very good lot. I was particularly impressed with the quality of the science and both the abundance and enthusiasm of young ornithologists in attendance. The three presentations by winners of our early professional awards—Michael Butler, Richard Ton, and Nancy Chen—were absolutely outstanding, for example. I sat in front of a well-known geneticist who said ‘wow’ twice and gasped once during Nancy Chen’s talk!

It would not be much of a stretch to predict that the 2018 meeting in Tucson will also be excellent.

This slideshow requires JavaScript.

SOURCES

  • Ahlquist JE (1999) Charles G. Sibley: A commentary on 30 years of collaboration. Auk 116:856–860.

  • Brush A (2017) François Vuilleumier, 1938–2017. The Auk 134: 776-777.
  • Clutton-Brock T, Ridley A (2017) Obituary: Amotz Zahavi 1928–2017, Behavioral Ecology 28: 1195–1197, https://doi.org/10.1093/beheco/arx115
  • Davies N (2018), Sir Patrick Bateson 1938–2017. Ibis 160: 253–254. doi:10.1111/ibi.12550

  • James FC (2017), Chandler Seymour Robbins (1918–2017). Ibis 159: 940–941. doi:10.1111/ibi.12518
  • Sheppard JM, Dawson DK, and Sauer JR (2017) Chandler S. Robbins, 1918–2017. The Auk 134: 935-938.

IMAGES: all of the images in this slideshow are in the public domain, or in the archives of the AOS History Committee. They are used here for educational purposes and may not be copied or used without permission from Jen Owen and/or the AOS History Committee.

Callow Youth

BY: Bob Montgomerie, Queen’s University | 11 December 2017

In science, controversies often arise over complex issues when researchers approach a problem from different points of view, backgrounds, and philosophies—think, for example, of the debates over nature vs nurture, selection vs drift, group vs kin vs individual selection, gradualism vs punctuated equilibria, and mechanisms of sexual selection. As Ledyard Stebbins pointed out in 1982, the resolution of these controversies often settles somewhere in the middle because, in fact, both sides were correct to some extent [1]. Usually scientific controversies churn away towards some resolution with some degree of civility, at least in public forums. But not always…

I can still remember my first experience with a less-than-polite public argument among ornithologists. This occurred during a talk at an AOU meeting in the 1970s when two prominent systematists got into a shouting match during the question period after a graduate student’s talk. The issue was whether the approach taken by the student was correct, and the argument seemed to be between the cladists and the pheneticists.

I learned later that these clashes among systematists were commonplace in the 1960s and early 70s, during a period that was later referred to as the ‘Systematics Wars’ [2]. I was doubly surprised at such ad hominem attacks because the systematists that I knew from my years as a young volunteer and employee at the Royal Ontario Museum—L.L. Snyder, Jim Baillie, Jon Barlow, Jim Rising, and Allan Baker—seemed like the kindest of men. My own field of behavioural ecology, while often dealing in controversy, does not seem to have descended into the sort of personal attacks that characterized those arguments about systematics in the mid 1900s.

While scouring some older literature, we recently discovered [3] that the ‘Systematics Wars’ period was not the first time that avian systematists had engaged in rather nasty exchanges. During the 1830s, for example, two young  brothers—Charles Thorold Wood and Neville Wood, tried to change the rules about naming birds in a way that involved heated debate, vitriol, and ad hominem attacks in their publications.

The Wood boys were wealthy, aristocratic, well-educated, and precocious. In 1835, when he was only 18, Charles published a quirky book—The Ornithological Guide—comprising 236 pages of poetry about birds, a compendium of ornithological books, each briefly reviewed, and a catalog (list) of the birds of Britain. Not to be outdone by his older brother, Neville published two books the next year, when he turned 18—British Song Birds and Ornithologists’ Text-book. British Song Birds alone ran to 400 pages, and, though it appeared not to include any novel observations, summarized much of what was then known about each species. Notably, none of the Woods’ publications included any illustrations.

The Woods felt that the names of birds—both scientific and English—were in a chaotic state and needed rules to make them more ‘scientific’. They were right. Even though Linnaeus had proposed his binomial system almost a century earlier [4], rules for zoological nomenclature that would provide some consistent structure and process were just beginning to be proposed in the 1830s [5].

GOULDbullfinch1837
Female and male European Bullfinches, by John Gould 1837 in his Birds of Europe

The English names were more confusing and disorganized. In England, some birds were even given different English names in different parts of the country. In the 1800s, for example the European Bullfinch was variously called: Bull Flinch (Yorks), Bull Head, Bulldog, Bull Spink, Bully (Yorks), Thick Bill (Lancs), Alpe, Hoop, Hope (SW), Tope, Hoof, Cock Hoop (Hereford), Olf (E Suffolk), Nope (Staffs/Shrops), Mwope (Dorset), Mawp (Lancs), Pope (Dorset), Red Hoop (m, Dorset), Blood Olp (m, Surrey/Norfolk), Tawny (f, Somerset), Tony Hoop, Tonnihood (f, Somerset), Black Cap (Lincs), Billy Black Cap, Black Nob (Shrops), Monk, Bud Bird, Bud Finch, Bud Picker (Devon), Budding Bird (Hereford), Plum Bird, and Lum Budder (Shrops) [6].

The Woods were sure that some logic and rules for the structure of English names were needed if ornithology was to develop as a science. As Neville said in one paper ”“If the proper English generic names were applied to every bird, how greatly would the acquisition of this fascinating study be facilitated! . . . By using the names which I have given above, this [confusion] is remedied, and all becomes plain and easy to understand.” [7] Their solution, for English names, was to make every bird within a genus have the same ’surname’, just as every genus had the same first name in the Linnaean system of Latin names. Thus, for example, Neville proposed:

Analyst1

It was also suggested  at the time [8] that English names be cleansed of modifiers that indicate size, abundance, location, or honorific. Thus present-day names like Little Cuckoo, Common Cuckoo, Moluccan Cuckoo, and Klaas’s Cuckoo would all have to be changed in the Cuculidae alone. Fortunately, cooler heads prevailed, as Hugh Strickland argued that common names were “consecrated by usage as much as any other part of the English language”. As he noted “the science of ornithology does not suffer by this incorrect application of English names, because those familiar appellations have no real or necessary connection with science”. Despite his admonition, the English names of birds are still in a state of flux today and may always be.

The Woods’ proposals were well reasoned, if sometimes seeming to be a little bizarre today. But their suggestions were often presented in such a way that demeaned those who had different points of view. In the Preface to his Bird Song Birds, Neville says: “while I agree with my predecessors in many points, I have found much to correct, and still more to add, to the meagre and unsatisfactory accounts of most of our British Ornithologists.” Ouch!

Neville’s review of Eleazar Albin’s Natural History of Birds claimed that it was “of no use in the present day” and of Jennings Ornithologia that he ”never had the misfortune to meet with a book so full of errors . . . We should have considered such a work beneath our notice, as it is impossible it can have the smallest connection with the advancement of Ornithology”. [9] Their writings are sprinkled with derogatory epithets and harsh criticism that even today strikes me as ungentlemanly, possibly simply the result of the Woods being callow youth [10].

Possibly because they gained few converts, and appear not have been well liked, both Charles and Neville drifted off to other pursuits before their 35th birthdays, never again to write about birds or to heap opprobrium on their fellow ornithologists. Charles published his last paper on birds (again on nomenclature) when he was 19 and then seems to have vanished from the public record; Neville trained as a doctor, and at the age of 26 moved to London where he practiced the new medical system called ‘homeopathy’ for the rest of his life. At least if he had stayed with birds he would have done something potentially useful.

Stebbins ended his 1982 address to the American Association for the Advancement of Science [11] by saying that: “My final hope is that evolutionists having different backgrounds and viewpoints will reduce their rivalry and collaborate increasingly in zealous research toward finding answers to these and other questions of major significance.” This was a noble sentiment but ignored those ineluctable human (or at least male) foibles of ego, ambition, and competitiveness. I would argue that controversies are are valuable part of scientific progress, and I have certainly enjoyed watching the systematics wars from the sidelines. We should always try to maintain civility but the controversies that arise from different points of view often move a field forward faster and more profitably than would the absence of skepticism about published research.

SOURCES

  • Albin E (1731) A Natural History of Birds : Illustrated With a Hundred and One Copper Plates, Curiously Engraven From the Life. Vol I. London: Printed for the author and sold by William Innys in St. Paul’s Church yard, John Clarke under the Royal-Exchange, Cornhill, and John Brindley at the King’s Arms in New Bond-Street. <available here>

  • Anonymous “N. F.” (1835) Remarks on vernacular and scientific ornithological literature. The Analyst 2: 305–307.

  • Birkhead TR, Montgomerie R (2016) A vile passion for altering names: the contributions of Charles Thorold Wood jun. and Neville Wood to ornithology in the 1830s. Archives of Natural History 43:221–236.

  • Greenoak F (1997) British Birds: their folklore, names, and literature. London: Christopher Helm.

  • Jennings J (1829) Ornithologia, or, the Birds: A Poem, in Two Parts : With an Introduction to Their Natural History; and Copious Notes. London: Sherwood, Gilbert, and Piper. <available here>

  • Stebbins GL. (1982a). Modal themes: a new framework for evolutionary synthesis. in Milkman R (Ed.), Perspectives on Evolution (pp. 12-14). Sunderland, MA: Sinauer Associates.
  • Stebbins GL (1982b) Perspectives in evolutionary theory. Evolution 36:1109–1118.
  • Sterner B, Lidgard S (2017) Moving past the Systematics Wars. Journal of the History of Biology https://doi.org/10.1007/s10739-017-9471-1
  • Wood CT (1835) The ornithological guide: in which are discussed several interesting points in ornithology. London Whittaker. <available here>
  • Wood N (1836a) British song birds: being popular descriptions and anecdotes of the choristers of the groves. London: John W. Parker. <available here>
  • WOOD N (1836b) The ornithologists’ text-book: being reviews of ornithological works on various topics of interest. London: John W. Parker. <available here>

Footnotes

  1. Stebbins (1982a) argued that modal themes were often the correct resolution of these conflicts, as has proven to be more-or-less the case in all the controversies listed here, with the notable exception that group selection is still controversial (and mistaken in my opinion)
  2. see, for example, Sterner and Lidgard (2017) for an overview
  3. Tim Birkhead made this discovery while researching for the project on Francis Willughby that he wrote here about a few weeks ago
  4. Linnaeus Systema Naturae had built on Willughby and Ray’s attempt at organizing and logically giving each species a scientific name (Ray (1678), but in the subsequent century there were many attempts to apply Linnaeus system to the birds with each author claiming authority and none of today’s rules about priority, coordination or homonymy.
  5. The International Zoological Congresses of 1889 and 1892 saw the first discussions about some universal rules in zoology but the International Rules on Zoological Nomenclature were not published until 1905.  Earlier, the ornithologist Hugh Strickland had published a set of 22 rules for the formulation of scientific names that formed some of the basis for this later code.
  6. the sex that was so named,  and the county or general area where each name was used, are shown in brackets, from Greenoak (1997).
  7. quotation from Wood 1835a: 238
  8. this was proposed in a paper in The Analyst signed simply N.F. (Anonymous 1835), who I think may well have been Neville Wood based on what N.F. says and the style of writing.
  9. see Birkhead and Montgomerie (2016) for some other gems. The paper is behind a paywall but send me an email if you cannot get it and would like a copy.
  10. ‘callow’ seems to me to be a particularly good adjective to describe these young men, as it refers to someone who is “inexperienced and immature”,  but in the 17th century the term was applied to birds, and meant ‘without feathers’ or immature and not ready to fly.
  11. presented on 7 Jan 1982, at a AAAS symposium marking the 100th anniversary of the death of Charles Darwin; quotation from Stebbins (1982b)

Pigeon Coup

BY: Bob Montgomerie, Queen’s University | 27 November 2017

When I was a young teenager I spent my Saturday mornings during the school year at the Royal Ontario Museum (ROM). I was there to attend the weekly meeting of the Toronto Junior Field Naturalists’ Club, but often stayed afterward to explore the public galleries. I particularly loved the dioramas of birds and mammals as they took me to distant places and bygone times that I could only dream or read about. In those days, there were virtually no nature documentaries on TV and precious few in the theatres [1].

Of all those superb dioramas, my favourite was the Passenger Pigeon, showing immense flocks (painted) descending into a forest clearing scattered with pigeons (mounted specimens) foraging on the acorns:

The scene reproduced depicts an April morning in the 1860s near Forks of Credit, Ontario…The visitor inspecting the exhibit should imagine himself standing at the edge of an old beech-maple forest overlooking the pioneer’s clearing. The scene is as we might have found it in the 1860s. The great pigeon flight is underway and will perhaps continue throughout the day. [2]

That diorama was opened to the public in 1935, the brainchild of Lester L. Snyder, the curator of birds, and constructed and painted by E. B. S. Logier who had joined the museum as illustrator in 1915. My memories of those dioramas came flooding back a couple of weeks ago when I saw the names Mark Peck and Allan Baker [3]—both from the ROM—among the authors of a new paper out of Berth Shapiro’s lab (UC Santa Cruz) on Passenger Pigeons published in Science, but more on that in a minute.

13662-Passenger Pigeon-retouched
Painted backdrop of the Passenger Pigeon diorama at the ROM (1935-1981)

Standing in front of that diorama I can remember thinking that a bird that had once been that abundant could not possibly be extinct. After all, Peterson’s Field Guide still illustrated them (in  a head and shoulders vignette on p 181 of my copy) so maybe he thought the bird might still be seen. My teen birding buddies and I spent many an afternoon naively scouring flocks of Mourning Doves just in case. After all, bison were also once extremely abundant, and hunted relentlessly, but were still round, albeit in small numbers.

I was also heartened by the fact that even if the species was really extinct, there must be thousands of specimens in museum collections that could be used for further study, since the ROM alone appeared to have so many that they could fill a diorama with mounted specimens alone. When I mentioned this to my friend and mentor Jim Baillie, assistant curator of birds at the ROM, he just laughed and told me there were only about 1500 specimens worldwide, of a species that once numbered in the billions. The reason for this wealth of specimens at the ROM, he said, was that they had been the beneficiaries of what he considered to be a major coup, when a local naturalist, musician and businessman [4], Paul Hahn, had decided—shortly after the Passenger Pigeon went extinct in 1914—to donate to the ROM as many specimens as he could locate, as a way “to ensure that future generations would know at least how handsome a bird it was.” [5]

thumbs_our-team_02
Paul Hahn

Hahn was born in Germany in 1875 but moved to Toronto with his family in 1898. In 1902 he saw his first Passenger Pigeon, a mounted specimen in a farmhouse north of the city and decided then to “set about gathering as many as possible of the mounted birds scattered around the country, both for the sake of future students and with the intention of preserving at least some specimens of a bird that would probably soon be extinct.” [5]. He presented his first specimen to the ROM in 1918 and had donated 70 by the time he died in 1962.

As a result of Hahn’s generosity, the ROM had 124 Passenger Pigeon skins and mounts by 1962, more than any other collection worldwide. I know this because in 1957 Mr Hahn started compiling a list of all the specimens of 7 extinct (our nearly so) bird species [6] held in museums and private collections around the world. Hahn died before his list could be published but Baillie took up the task, seeing it through to print in 1963 as a book Where is that Vanished Bird? That book lists every specimen (including skeletons) known to Hahn [7], its date and place of collection, its sex, the collector, and the current collection in which it was held.

ROM2013_13563_1
One of the Passenger Pigeon mounts at the Royal Ontario Museum

The first Passenger Pigeon specimen whose collection date was known was a male taken in the Carolinas in about 1810, housed with one other specimen (a female, from Georgia collected in 1821) in the Zooligische Museum in Berlin. The Naumann Museum in Köthen that Tim Birkhead wrote about last week also had a male, collected in about 1830 (locality unknown).

The recent Science paper made good use of the ROM collection of Passenger Pigeons, analyzing the DNA extracted from the toe pads of 84 specimens, 63 of which were from the ROM. Analyzing both nuclear and mitochondrial genes, the researchers confirmed that the Passenger Pigeon had surprisingly low genetic diversity. This low diversity is unexpected because large populations are predicted from theory to be genetically diverse, and you don’t get larger bird populations than those of the Passenger Pigeon. To explain this loss of diversity, the authors argued that it was driven by high rates of dispersal and adaptive evolution that removed harmful mutations. Such low diversity would have made the species particularly susceptible to disease or environmental change, two factors that might have doomed the species once populations had been decimated by hunting. This study also concluded, based on some sophisticated genomic analyses, that Passenger Pigeon populations had probably persisted at extremely high numbers for 20,00 years or more before the 1800s [8].

The Passenger Pigeon diorama at the ROM was dismantled in 1981, in part because it was showing its age, but also because the age of dioramas was over, replaced in part by the ubiquitous nature shows in TV. That saddens me but I am more than ever convinced that clubs for young field naturalists, and museums that store and preserve specimens, deserve our unending support.

SOURCES

  • Hahn P (1963) Where is that Vanished Bird? Toronto: University of Toronto Press.

  • Hung C-M, Shaner P-JL, Zink RM, Liu W-C, Chu T-C, Hiuang W-S et al. (2014) Drastic population fluctuations explain the rapid extinction of the passenger pigeon. Proceedings of the National Academy of Scienes USA 111:10636–10641.

  • Murray GGR, Soares AER, Novac BJ, Schaefer NK, Cahill JA, Baker AJ et al. (2017) Natural selection shaped the rise and fall of passenger pigeon genomic diversity. Science 358:951–954.


FOOTNOTES

1. The first nature documentaries on TV were a series called Fur and Feathers on Canadian Broadcasting Corporation (CBC) channel in 1955-56, in black and white (of course). By 1960, Disney had produced 14 movies in its True-Life Adventures series, including the The Living Desert, The Secrets of Life, African Lion, and White Wilderness all of which enthralled my naturalist friends and I when they played at our local theatre.

2. Text from the ROM’s Passenger Pigeon diorama, courtesy Mark Peck, 22 Nov 2017.

3. Mark Peck is Ornithology Technician at the ROM, where Allan Baker (1943-2014) worked for 42 years as both a curator of birds and eventually head of their Department of Natural History.

4. Hahn was an accomplished cellist who played with the Toronto Symphony Orchestra. He also founded Paul Hahn Pianos in Toronto in 1913, a company that is still in business today.

5. Quotations from Hahn 1963:1.

6. As of 1962: skins and mounts of 1532 Passenger Pigeons, 365 Eskimo Curlews, 78 Great Auks, 720 Carolina Parakeets, 413 Ivory-billed Woodpeckers, 54 Labrador Ducks and 309 Whooping Cranes (Hahn 1963).

7. Beginning in 1957 he sent out questionnaires to people and museums that he thought might know or know about those specimens. He got more than 1000 response.

8. Based on DNA samples from only 3 Passenger Pigeons, Hung et al. (2014) performed a different genomic analysis and concluded that population sizes had fluctuated dramatically—only occasionally reaching numbers in the billions—thereby increasing its risk of extinction during population lows. Evaluating the conclusions of these two studies is above my pay grade but I expect that both labs will argue that their analyses are correct.

IMAGES: ROM photos by Brian Boyle, courtesy of Mark Peck (both at the ROM); Paul Hahn from the Paul Hahn & Co. website at https://paulhahn.com/about/who-we-are/